AOU Classification Committee – North and Middle America Proposal Set 2017-B No. Page Title 01 02 Recognize additional species in the Aulacorhynchus “prasinus” toucanet complex 02 17 Treat the subspecies (A) spectabilis and (B) viridiceps as separate species from Eugenes fulgens (Magnificent Hummingbird) 03 23 Elevate Turdus rufopalliatus graysoni to species rank 04 26 Recognize newly described species Arremon kuehnerii 05 30 Revise the classification of the Icteridae: (A) add seven subfamilies; (B) split Leistes from Sturnella; (C) resurrect Ptiloxena for Dives atroviolaceus; and (D) modify the linear sequence of genera 06 34 Revise familial limits and the linear sequence of families within the nine- primaried oscines 07 42 Lump Acanthis flammea and Acanthis hornemanni into a single species 08 48 Split Lanius excubitor into two or more species 09 54 Add Mangrove Rail Rallus longirostris to the main list 10 56 Revise the generic classification and linear sequence of Anas 1 2017-B-1 N&MA Classification Committee p. Recognize additional species in the Aulacorhynchus “prasinus” toucanet complex Background: The AOU (1998) presently considers there to be just one species of Aulacorhynchus prasinus, which ranges from Mexico to Guyana and Bolivia. This taxon’s range combines the taxonomic oversight regions of both the North American and South American classification committees, so this proposal is designed to be submitted to both, with committee-structured voting sections at the end. This is easy to do biologically, because the taxa fall out fairly neatly split between North and South America. (The Panamanian blue-throated population breeding on Cerro Tacarcuna (subspecies cognatus) has (Hilty and Brown 1986) and has not been (Donegan et al. 2015) included in the Colombian avifauna.) The AOU’s first treatment of this group in Middle America began with the geographic expansion undertaken in the sixth edition of the Check-list (AOU 1983). The historic treatments of the genus are given in Table 1 (from Winker 2016). In brief, evidence of hybridization caused massive lumping into a broadly defined prasinus from Peters (1948) onward, with recent genetic evidence of divergence causing some authors to propose that the prasinus complex is made up of as many as seven species (Table 1). These recent proposals have not been widely accepted; I summed the situation up as follows (Winker 2016): “Renewed interest in this complex (Navarro et al., 2001; Puebla-Olivares et al., 2008; Bonaccorso et al., 2011; Del Hoyo & Collar, 2014) is beginning to rectify the absence of data, but the ensuing taxonomic changes recommended have either been based on a different species concept (Bonaccorso et al., 2011) or have inadequately considered the hybridization and intergradation (e.g., Navarro et al., 2001; Puebla-Olivares et al., 2008; Del Hoyo & Collar, 2014) that have been integral to supporting the “post-Peters” taxonomy. These latter works have recommended elevation of numerous A. prasinus (sensu lato) taxa to species status (Table 1), but they did not address the reasons for lumping in the first place: evidence of hybridization. There has also been heavy reliance on a single molecular marker (mtDNA) for species delimitation in the A. prasinus complex (Puebla-Olivares et al., 2008; Bonaccorso et al., 2011). This is problematic because mtDNA can be misleading about species limits and relationships between populations due to gene-tree/species-tree mismatches and because genetic distance is not a reliable indicator of species limits (Avise & Wollenberg, 1997; Irwin, 2002; Funk & Omland, 2003; Degnan & Rosenberg, 2006; Cheviron & Brumfield, 2009; Galtier et al., 2009; Ribeiro, Lloyd & Bowie, 2011; Toews & Brelsford, 2012; Pavlova et al., 2013; Peters et al., 2014; Dolman & Joseph, 2015; Morales et al., 2015). Thus, species limits in the group remain uncertain (Table 1).” 2 There are six color-based groups in the prasinus complex, within which some have additional described subspecies. These major groups have been recognized through much of the history of the taxon (Table 1) and were reaffirmed by the analyses of del Hoyo and Collar (2014). The characters upon which they are based are given in Winker (2016: table 2) and can be seen in the accompanying Plate. Figure 1. The six major, color- based taxonomic groups of the Aulacorhynchus “prasinus” species complex, from top to bottom: A) wagleri; B) prasinus (nominate prasinus and warneri, the full-bodied bird, are portrayed): C) caeruleogularis; D) albivitta (griseigularis and nominate albivitta are portrayed); E) cyanolaemus (yellow-tipped bill); and F) atrogularis. 3 Table 1. Treatments of species-level diversity in the genus Aulacorhynchus. Taxa historically recognized only as subspecies are not included (see text for these taxa in "prasinus"). An X means the taxon was treated as a species, a dash indicates not available to be treated yet, and a blank indicates that the taxon was not considered. Nav. et al. (2001)g S & G P-O et al. Sclater (1896)a Sibley & Short & (2008)g Dickinson & del Hoyo & B & C Monroe Remsen (1891) (1912)b Cory (1919) Peters (1948) (1990) Horne (2001) B. et al. (2011)g (2013)i Collar (2014) Winker (2016) A. sulcatus × × × × × × × × × ssp. of × × A. erythrognathus ssp. of sulcatus ssp. of sulcatus ssp. of sulcatus ssp. of sulcatus sulcatus ssp. of sulcatus ssp. of ssp. of × × × × × A. calorhynchus sulcatus ssp. of sulcatus ssp. of sulcatus sulcatus A. derbianus × × × × × × × × × ssp. of ssp. of × × × × × × A. whitelianus derbianus derbianus A. haematopygus × × × × × × × × × A. coeruleicinctis × × × × × × × × × A. huallagae – c – c – c × × × × × × A. prasinus × × × × × × × × × × ssp. of × × × × × × A. wagleri ssp. of prasinus ssp. of prasinus prasinus ssp. of ssp. of × × × × × × A. caeruleogularis ssp. of prasinus prasinus ssp. of prasinus prasinus ssp. of ssp. of ssp. of ssp. of × A. cognatus – d – d caeruleogularis ssp. of prasinus ssp. of prasinus prasinus caeruleogularis caeruleogularis ssp. of × × × × × × A. albivitta ssp. of prasinus ssp. of prasinus prasinus ssp. of × A. griseigularis – e – e – e ssp. of prasinus ssp. of prasinus prasinus ssp. of albivitta ssp. of albivitta ssp. of × × × h A. lautus – f ssp. of prasinus ssp. of prasinus prasinus ssp. of albivitta ssp. of albivitta ssp. of ssp. of × × × × A. cyanolaemus ssp. of prasinus ssp. of prasinus atrogularis prasinus ssp. of atrogularis ssp. of ssp. of × × × A. dimidiatus ssp. of prasinus ssp. of prasinus atrogularis prasinus ssp. of atrogularis ssp. of atrogularis ssp. of × × × × × × A. atrogularis ssp. of prasinus ssp. of prasinus prasinus a – Salvin & Godman (1896) treated only Middle American Aulacorhynchus, which at the time were considered Aulacorhamphus. b – Brabourne and Chubb (1912) treated South American members of the genus (then considered Aulacorhamphus. c – huallagae was described by Carriker (1933). d – cognatus was described as a subspecies by Nelson (1912). e – griseigularis was described as a subspecies by Chapman (1915). 4 f – lautus was described by Bangs (1898). g – Navarro et al. (2001), Puebla-Olivares et al. (2008), & Bonaccorso et al. (2011) together included most Middle American and South American Aulacorhynchus taxa. h – though not included in either study. i – Treatment matches the South American Classification Committee (Remsen et al. 2016). 5 New Information: In Winker (2016) I tested the hypothesis that these are “cookie-cutter” (i.e., morphologically nearly identical) toucanets differing mostly in coloration. I also examined specimens carefully for phenotypic evidence of hybridization. A couple of key factors were central to my treatment of the group. First, these birds move about considerably during the nonbreeding season, providing hypothetical opportunities for gene flow across zones of nearest approach. “For example, in south- central Mexico (Oaxaca), A. prasinus and A. wagleri breed within about 100 km of each other, a distance that A. prasinus individuals appear to move routinely away from their breeding areas, e.g., at the base of the Yucatan Peninsula (e.g., Land, 1970; Jones, 2003), which does not seem unusual for an arboreal frugivore (see also discussions in O’Neill & Gardner, 1974, and Navarro et al., 2001).” (Winker 2016). The hitherto unrecognized (although published by Puebla-Olivares et al. 2008) gene flow between albivitta and atrogularis in NE Ecuador indicates that this hypothesis has merit. Second, I considered that the likelihood of successful gene flow/reticulation between two lineages decreases with increased anagenesis or adaptive divergence, arguing as follows (Winker 2016): “Effective lineage reticulation requires that hybrid offspring have equal or greater fitness than offspring of pure parental forms. Also, gene flow must occur frequently enough to overcome the differentiating selective factors likely to be operating on largely allopatric populations (and this relationship is nonlinear; see Winker, 2010 for discussion). The more differences there are between populations in morphology, the more differences there are likely to be in selective factors operating on these populations and the more difficult effective gene flow is likely to be between populations; at larger scales this results in the general correlation between morphological difference and reproductive isolation (Mayr, 1963; Price, 2008).” Another important factor that I considered that did not seem to have been adequately addressed before is that named subspecies in this group do not represent equivalent